The Theropod Ancestry of Birds: New Evidence from the Late Cretaceous of Madagascar

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Abstract
A partial skeleton of a primitive bird, Rahona ostromi, gen. et sp. nov., has been discovered from the Late Cretaceous of Madagascar. This specimen, although exhibiting avian features such as a reversed hallux and ulnar papillae, retains characteristics that indicate a theropod ancestry, including a pubic foot and hyposphene-hypantra vertebral articulations. Rahona has a robust, hyperextendible second digit on the hind foot that terminates in a sicklelike claw, a unique characteristic of the theropod groups Troodontidae and Dromaeosauridae. A phylogenetic analysis placesRahona with Archaeopteryx, makingRahona one of the most primitive birds yet discovered.